VON HOLST, ERICH

(b. Riga, Latvia, 28 November 1908;

d. Herrsching, Bavaria, Germany, 26 May 1962), experimental zoology, ethology, physiology, musical instruments.

Von Holst is remembered for his outstanding experimental studies of the central nervous system, the flight of birds, and many other physiological problems. He was one of the founders of behavioral physiology and helped establish ethology as a scientific discipline. He was also a gifted musician and contributed to a scientific understanding of musical instruments.

Life and Passion. Von Holst, son of psychiatrist Dr. Walther von Holst and Dorothea von Holst, was born in Riga, Latvia, but attended school in Danzig (now Gdańsk, Poland). Serious health problems (articular rheumatism) in his childhood led to an incurable cardiac defect and thus negatively affected his life expectancy. He was constantly aware that his life might be rather short. However, early in his youth he discovered both his passion for animals and his enthusiasm for engineering. While recovering in a hospital, he read Alfred Brehm’s popular Tierleben—a comprehensive (in many aspects anthropomorphic) description of animal life—and was fascinated. Later he combined zoological phenomena with engineering to arrive at highly original solutions of physiological problems. Also, he developed a passion for music and musical instruments, particularly the viola. He was concerned with the construction of this instrument and the generation of its sound pattern.

His former students and colleagues describe von Holst as an outstanding personality with high moral standards, at the same time generous and helpful. He was an indefatigable and keen worker, eager to solve scientific problems. Neither in his professional nor in his private life was he willing to make—or to accept—a compromise. He lived very intensively, due to his early recognition of his heart disease. Worry about death was combined in him with an enormous energy and a tremendous drive to live and to work. The well-known philosophy of “live for today, for there may be no tomorrow” could very well apply to him. This can also help explain the variety of his interests and the remarkable scientific work that he accomplished within just a few decades. His students were allowed to contact him and ask for help at almost any hour of the day (or night), yet he was rigorous when it came to scientific work and did not accept any indolence. Passionate students with original ideas found in him a benevolent supporter. But he could be also merciless in his criticism. In discussions after a lecture it frequently happened that he sharply attacked and discouraged the lecturer. Needless to say, in the eyes of some people he was a difficult man.

Early Scientific Work. Von Holst studied in Kiel, Vienna, and Berlin. Already in his doctoral thesis, which dealt with the function of the nervous system in earthworms (and was published in 1932 in a well-recognized zoological yearbook), he found arguments against the doctrine according to which the behavior of organisms is governed by reflexes. In many experiments he found out that the locomotion of earthworms is not caused by a chain of reflexes stimulating one segment after the other, but that excitations in the head autonomously—without any further stimuli—affect the other parts of the animal.

To appreciate this achievement, one has to remember the philosophical view guiding the study of behavior at that time. In the 1930s the majority of physiologists and behavior scientists were convinced that any organism’s activity is mainly a reflex action and that the organism is initially a clean slate or tabula rasa (behaviorism). They followed the tradition of the Russian physiologist Ivan P. Pavlov and defined behavior essentially as the response of living beings to some kind of (external) stimulus (empty-organism doctrine). Thus, they disregarded any notion of innate or instinctive behavior and accepted only those behavioral traits as relevant that could be studied and manipulated in experiments. On the other side, there was a smaller faction of biologists and psychologists holding that some “inner mechanisms” were the very driving forces of animal behavior (purposive psychology, Zweckpsychologie).

The opposition between these two schools reflected pretty well the old battle between mechanists and vitalists, which had been ignited by the venerable question how, after all, living beings and their properties are to be explained: as mere machines or complex systems exhibiting some “inner” forces and purposes qualitatively different from physical laws. The question is now obsolete for it is known that any living system is not just a machine but that its specific properties can be sufficiently explained without resort to cryptic vital forces and the like (which, anyway, have no explanatory power). In the early 1930s, however, the methodological stance of biology was different from its status in the early twenty-first century.

Thus, when von Holst entered the scene, the situation in the behavior sciences was somewhat confusing. Von Holst was not at all inclined to vitalism, at the same time he did not embrace the doctrine that the study behavior is to be reduced to the study of reflexes. His own studies were pathbreaking and gave behavioral physiology a new direction. As a neurophysiologist he regarded nervous actions as somehow autonomous and became

particularly interested in the spontaneous action of the central nervous system in addition to the aspect previously so widely studied: the relationship between stimulus and response. He was able to demonstrate that the central system does not passively link stimuli to responses but is highly active in itself, producing “internalstimuli” and incorporating numerous functional systems. (Thorpe, 1979, p. 110)

Von Holst’s early scientific work already gave some impetus to the emerging field of ethology, the comparative study of behavior from a biological perspective. Other leading figures in the history of this field are, above all, Oskar Heinroth, Karl von Frisch, Konrad Lorenz, and Nikolaas Tinbergen. Later, due to his many outstanding experimental studies, von Holst would be regarded as one of the founders of behavioral physiology, the discipline concerned with the physiological basis of all kinds of animal (and human) behavior. He introduced the cybernetic approach to ethology and showed that behavioral phenomena are not simply caused by outer stimuli but are rather to be understood on the basis of complex feedback and regulatory principles. In other words: A living being does not simply react to its surroundings, it is not—as the empty-organism doctrine of the behaviorists suggested— “molded” by its environment, but is an active system searching for its possibilities to live and survive.

Professional Career. After he had obtained his PhD, von Holst worked as a fellow in Frankfurt am Main, where he was attracted by the approach of the physiologist Albrecht Bethe, whose conception of plasticity (Plastizitätslehre) clearly contrasted with the doctrine of reflexes. In Bethe he found the ideal of an experimental biologist. He combined Bethe’s methodology with his own findings from his doctoral thesis. In 1934 and 1935 von Holst took the position of an assistant at the Zoological Station in Naples (Stazione Zoologica di Napoli). From 1936 to 1938 he worked in Berlin and finished his Habiltation, a physiological study on the spinal cord. After that, he worked for some years as an academic teacher without salary (privatdozent) in Göttingen, and in 1946 he was appointed full professor of zoology at the University of Heidelberg. Immediately after World War II, von Holst became cofounder and coeditor of the Göttinger Universitätszeitung, a journal that expressed the new intellectual freedom after the Nazi regime.

From 1948 on he was head (Abteilungsleiter) at the Max Planck Institute for Marine Sciences in Wilhelmshaven (Lower Saxony), where, in 1950, he organized a workshop in ethology that was the prelude, so to speak, to many other regularly organized ethological symposia in the following years. One of the participants of the workshop, W. H. Thorpe, described the conditions in Wilhelmshaven as “curious and inconvenient.” The institute was housed in an unfinished naval barracks building; the quarters were cramped, and the staff and students lived closely packed. Nevertheless, in retrospect, this was the important beginning of “institutionalized ethology.” Von Holst was eager to establish his discipline, behavioral physiology—and he was very successful. His idea of a research group combining different approaches to the study of behavior was realized in the 1950s when the Max Planck Society approved an institute in Seewiesen (Bavaria) that soon became one of the world’s leading institutes in the field.

The Max-Planck-Institut für Verhaltensphysiologie (behavioral physiology) was inaugurated in 1958. Von Holst was appointed director of the institute, Lorenz his codirector. The name of the institute paid tribute to von Holst’s work. In fact, his reputation at the Max Planck Society was most remarkable, his scientific integrity undisputed. Already in 1950 he was given a special role in one of the (still) leading scientific journals in Germany, Die Naturwissenschaften (the official organ of the Max Planck Society for the Advancement of Science). He became a kind of “super editor” and was asked to shape the direction of this periodical. Also von Holst was, for some decades, coeditor of the Zeitschrift für vergleichende Physiologie(Journal of comparative physiology).

The name of the institute in Bavaria (Verhaltensphysiologie) was somehow misleading. Lorenz with his research group was doing mainly descriptive and comparative ethology. Behavioral physiology was only one of the disciplines united in Seewiesen. After von Holst’s early death in 1962 the institute’s leading figure was definitely Lorenz. But by that time the physiological approach to the study of behavior was already well established. This is not to say that Lorenz’s and von Holst’s approaches contradicted each other. As Lorenz and von Holst had very different personalities, they advocated different views of the “philosophy” of behavior sciences. But each of them was needed. Brought together, they formed the whole corpus of ethology as a comprehensive study of behavior in animals as well as humans.

Further Scientific Work. In 1933, after his doctoral dissertation, von Holst dedicated his work to a variety of physiological problems and their solution. During his stay in Naples, he observed that some fish—under particular experimental constraints—do not coordinate their fins in an absolute manner, so to speak, although they do not move them separately, one independent from the other. He called this phenomenon “relative coordination” and— in a long series of experiments—found a relation between two centers in the central nervous system: an autonomous and a dependent one. Relative coordination produces a harmony between individual rhythms (e.g., in the fins of a fish). The better this can be achieved, the more stable is the end result: the coordination of the rhythms.

In other studies, von Holst was concerned with—if not obsessed by—the flight of animals, particularly birds. In the search to understand this phenomenon, he built flying objects and constantly rebuilt them so that, in the end, the results were models of flying animals exhibiting a certain autonomy. The intuition of a talented engineer and experimentalist also enabled him, for example, to construct a model of the flying Mesozoic reptile, Rhamphorhynchus. The participants of a paleontological meeting were impressed when his artificial pterosaur was flying in the conference room.

The physiology of the nervous system offered von Holst further topics for study. In 1950 his seminal paper “Das Reafferenzprinzip” was published (coauthored by his student Horst Mittelstaedt). The principle of reafference contradicts the stimulus-response conception of the behaviorists and describes mechanisms that check and recheck any newly acquired motor patterns. It is a regulatory principle in the nervous system to control the external stimulus.

In the following years von Holst published many other papers, most of them reporting the results of his own experimental studies that he always carried out with passion. He constantly devised new—and improved old—experimental methods to answer physiological questions. He offered most convincing evidence of the physiological autonomy (Eigenaktivität) of the central nervous system. This can be regarded his greatest achievement. Generally, his work clearly demonstrates the complex functionality of organisms and their activities that are not governed by external stimuli and reflexes. Von Holst contributed to a better understanding of the concept of instinct, a previously somewhat confusing and obscure concept.

In his experimental work he was dedicated to analytical thinking; however, he did not neglect the importance of synthesis and the “holistic” approach. In many of his writings, one can find general statements concerning the methodology and philosophy of science. He considered mere counting and measurement insufficient and claimed that truly scientific progress is achieved only through qualitatively new insight. Also, he argued that at the beginning of any physiological investigation the investigator should try to find him- or herself in the situation of the constructor of the respective animal. This is an interesting methodological proposition that at least in von Holst’s case led to remarkable and astonishing results.

It may also be noted that von Holst was a brilliant speaker. He did not read a manuscript, but gave in his talks the impression that he was developing his ideas just then. He always encouraged his students to give their presentations the same way. Because of his early death, many of his projects remained unfinished, yet his pioneering work and his meaning for the development of ethology as a particular branch of biology (with essential implications for other sciences, especially psychology) are appreciated by all students working in this field.

Finally, von Holst has to be remembered as an artist. He was a gifted musician. As a violist he could attract and fascinate his listeners for many hours. In earlier years, he used to say that the scientist and the artist are different aspects of his personality. Later, however, he successfully combined his enthusiasm for science with his passion for music and started to study musical instruments in a way similar to his studies of living beings. Several of his publications give evidence of his systematic inquiry into the construction, function, and sound pattern of the viola. In the history of music there are certainly not many people who combined their deep knowledge of physiological processes so successfully with an understanding of the architecture and function of musical instruments. Von Holst started to write a book, Geigenkunde für Liebhaber(Violin science for amateurs), which remained unfinished.

BIBLIOGRAPHY

WORKS BY VON HOLST

“Untersuchungen über die Funktion des Zentralnervensystems beim Regenwurm.” Zoologische Jahrbücher, Allgemeine Zoologie 51 (1932): 547–588; 53 (1932): 67–100.

“Studien über die Reflexe und Rhythmen beim Goldfisch (Carassius auratus).” Zeitschrift für vergleichende Physiologie 20 (1934): 582–599.

“Über den Prozess der zentralen Koordination.” Pflügers Archiv 236 (1935): 149–158.

“Versuche zur Theorie der relativen Koordination.” Pflügers Archiv 237 (1936): 93–121.

“Vom Wesen der Ordnung im Zentralnervensystem.” Naturwissenschaften 25 (1937): 625–631, 641–647.

“Über die nervöse Funktionsstruktur des rhythmisch tätigen Fischrückenmarks.” Pflügers Archiv 241 (1939): 569–611.

“Über die relative Koordination bei Arthropoden.” Pflügers Archiv 246 (1943): 847–865.

“Über ‘künstliche Vögel’ als Mittel zum Studium des Vogelflugs.” Journal für Ornithologie 91 (1943): 406–447.

“Vom Flug der Tiere und vom Menschenflug der Zukunft.” Schriften der Universität Heidelberg 3 (1948): 95–112.

“Von der Mathematik der nervösen Ordnungsleistungen.” Experientia 4 (1948): 374–381.

“Die Tätigkeit des Statolithenapparates der Wirbeltiere.” Naturwissenschaften 37 (1950): 265–272.

With H. Mittelstaedt. “Das Reafferenzprinzip.” Naturwissenschaften 37 (1950): 464–476.

“Ein neuer Vorschlag zur Lösung des Bratschenproblems.” Instrumentenbau-Zeitschrift 8 (1953): 46–48.

“Physiologie und Verhalten.” Mitteilungen der Max-Planck-Gesellschaft 5 (1954): 270–275.

“Relations between the Central Nervous System and the Peripheral Organs.” British Journal of Animal Behaviour 2 (1954): 89–94.

“Zentralnervensystem (Das Muskelspindelsystem der Säuger).” Fortschritte der Zoologie 10 (1956): 381–390.

“Aktive Leistungen der menschlichen Gesichtswahrnehmung.” Studium Generale 10 (1957): 231–243.

“Der Saurierflug.” Paläontologische Zeitschrift 31 (1957): 15–22.

“Wie flog Rhamphorhynchus?” Natur und Volk 87 (1957): 81–87.

With U. von Saint Paul. “Vom Wirkungsgefüge der Triebe.” Naturwissenschaften 47 (1960): 409–422.

Zur Verhaltensphysiologie bei Tieren und Menschen Edited by B. Hassenstein. 2 vols. Munich: Piper, 1969–1970. A collection of von Holst’s papers on behavioral physiology. Contains a list of obituaries.

OTHER SOURCES

Burkhardt, Richard W. Patterns of Behavior: Konrad Lorenz, Niko Tinbergen, and the Founding of Ethology. Chicago: University of Chicago Press, 2005.

Hassenstein, Bernhard. “Erich von Holst (1908–1962).” In Darwin & Co.: Eine Geschichte der Biologie in Portraits, vol. 2, edited by Ilse Jahn and Michael Schmitt. Munich: C. H. Beck, 2001.

Lorenz, Konrad. “Erich von Holst zum Todestag.” Die Naturwissenschaften 49 (1962): 385–386.

———. “Erich von Holst, Seher und Forscher.” 4. Biologisches Jahresheft des Verbandes deutscher Biologen (1964): 19–24.

Thorpe, W. H. The Origins and Rise of Ethology. London: Heinemann, 1979.

Wuketits, Franz M. Die Entdeckung des Verhaltens: Eine Geschichte der Verhaltensforschung. Darmstadt, Germany: Wissenschaftliche Buchgesellschaft, 1995.

Franz M. Wuketits